cetaceanCetaceaany member of an entirely aquatic group of mammals commonly known as whales, dolphins, and porpoises. The ancient Greeks recognized that cetaceans breathe air, give birth to live young, produce milk, and have hair—all features of mammals. Because of their body form, however, cetaceans were commonly grouped with the fishes. Cetaceans are entirely carnivorous, although members of the order Sirenia (manatees, dugongs, and Steller’s sea cow) were once referred to as the “herbivorous Cetacea.” In the past cetaceans were important resources (see whaling), but by the end of the 20th century their economic importance was almost solely due to whale watching, a tourist activity and major source of income for certain coastal regions of many countries.

Form and function

General features

Body surface

The hair covering that is common to mammals is drastically reduced in cetaceans, likely because hair is a poor insulator when wet and increases drag during swimming. Hairs on cetaceans are restricted to the head, with isolated follicles occurring on the lower jaw and the snout. These are thought to be remnants of sensory whiskers (vibrissae). External pigmentation is important to many animals as a basis for individual recognition and species recognition. Hair defines the colour pattern of most mammals, but, because cetaceans have very little hair, the outer layer of skin (epidermis) produces their markings, most commonly in shades of black and white. The appearance of some cetaceans is affected by various organisms living on or in the skin. Examples include yellow algae that colour the lower body surface of blue whales (Balaenoptera musculus) and the variety of whitish organisms living on bodies of gray whales (Eschrichtius robustus) and right whales (family Balaenidae).

Locomotor adaptations

The most noticeable adaptation of cetaceans to life in the water is their locomotive system. Because cetaceans descended from mammals that moved their limbs in a vertical plane rather than in a horizontal plane, they use vertical strokes when they swim, instead of horizontal strokes like a crocodile or fish. Cetaceans evolved from four-legged (quadruped) terrestrial animals, for which limbs played a primary role in movements, into virtually limbless aquatic creatures living in an environment where the back muscles are more important. Forelimbs are still present but are reduced to finlike flippers having shortened arm bones and no individual fingers. The hind limbs are lost entirely; only vestigial elements sometimes remain internally. Pelvic remnants occur in all cetacea but the dwarf and pygmy sperm whales. Flippers help to steer, while the back muscles, which are very large, drive the tail to propel the animal. Cetaceans have developed horizontal flukes that increase the propulsion area driven by the back muscles. Like fish, almost all cetaceans possess a dorsal fin that serves as a keel. The dorsal fin and flukes are composed of connective tissue, not bone. Other connective tissue, such as external ears, has been lost, and the male genitalia have moved internally.

Respiration

Normally, cetaceans breathe while moving through the water and spend only a short time at the surface, where they exhale in an explosive ventilation called a blow. The blow is expelled forcibly and can be compared to a cough. Cetaceans use up to 80 percent of their lung volume in a single breath, in contrast to humans, who use only 20 percent. The blow is visible because of water condensation and mucous particles; blows of blue whales are frequently more than 6 metres (20 feet) high. When a terrestrial mammal loses consciousness, it breathes reflexively, but breathing is not a reflex in cetaceans. Thus, when a cetacean loses consciousness, it does not breathe and quickly dies. For this reason, veterinarians had to perfect respirators before dolphins could be successfully anesthetized.

Circulation and thermoregulation

Cetaceans, like all mammals, have a four-chambered heart with paired ventricles and auricles. The pattern of circulation is similar to that of other mammals, with the exception of a series of well-developed reservoirs for oxygenated blood called the rete mirabile, for "marvelous network." These provide bypasses that enable cetaceans to isolate skeletal muscle circulation during diving while using the oxygen stored in the remaining blood to maintain the heart and brain—the two organs that depend on a constant supply of oxygen to survive.

Water conducts heat much more rapidly than air and is colder than the mammalian body temperature of about 37 °C (98.6 °F). Cetacean evolution has countered this problem in three ways: reducing external appendages that lose heat, developing an insulating layer of blubber, and developing countercurrent circulation to minimize heat loss. The reduction of various appendages as mentioned above also facilitates locomotion in water.

In whales, a layer of the skin (dermis) has evolved into a blanket of blubber, which is extremely rich in fats and oils and therefore conducts heat poorly. This blanket covers the entire body and is up to 30 cm (12 inches) thick in large whales, making up a significant portion of the animal’s weight. The oil yield of blubber from a blue whale, for example, was up to 50 tons.

The most important mechanism in cetacean thermoregulation is the development of countercurrent blood exchange, an adaptation that allows the animal to either conserve or dissipate heat as needed. Blood that drains from the surface of the skin has been cooled by close contact with the external environment, and it can return to the cetacean’s heart via two different routes. If it returns by the peripheral route, the blood courses back to the heart through superficial veins, where it continues to lose heat and arrives at the heart cool. This dumps the animal’s excess heat to the environment. Such heat shedding is particularly important to large whales because of their enormous surface area-to-volume ratio. If, however, the body temperature of the whale is already cool, the oxygen-depleted venous blood can instead return to the heart through vessels that are wrapped around arteries carrying warm blood to the periphery of the animal. Along this route the venous blood is warmed by the arterial blood and arrives at the heart warm. The arterial blood, having transferred its heat into the venous blood rather than the environment, arrives precooled at the surface of the skin.

Feeding adaptations

Before cetaceans evolved aquatic adaptations, they had a fully differentiated set of teeth (heterodont dentition), including incisors, canines, premolars, and molars. As the animals became more adapted to aquatic locomotion and lost the ability to manipulate food with their forelimbs, they started grabbing their food and swallowing it whole. In toothed whales (suborder Odontoceti), heterodont dentition declined and was replaced with a homodont dentition in which every tooth is a simple cone. The number of teeth varies among toothed whales, from two in the beaked whales (family Ziphiidae) to 242 in the La Plata river dolphin (Pontoporia blainvillei), to allow efficient capture of prey. Baleen whales (suborder Mysticeti), on the other hand, have lost all teeth in both jaws and instead have two rows of baleen plates in their upper jaws only. This apparatus enables baleen whales to consume vast quantities of small prey in a single mouthful.

In general, whales have relatively large mouths. The mouth of one adult bowhead, or Greenland right whale (Balaena mysticetus), measures five metres long and three metres wide and is the biggest oral cavity on record. The stomach in cetaceans is composed of four compartments: forestomach, main stomach, connecting chambers, and pyloric stomach. The forestomach is actually a dilation of the esophagus and is lined with simple epithelium (layers of flattened cells). It acts merely as a holding chamber and therefore is not a true stomach. The main stomach, lined with active gastric epithelium, is the first true digestive compartment, and it is followed by the small connecting chambers and the pyloric stomach. From there, food enters the small intestine through the pyloric sphincter and the duodenal ampulla. Most cetaceans do not have a cecum or appendix, and in most there is no anatomic difference between the small and the large intestine.

The senses

The sensory system of any animal can be divided into somesthetic senses—those relating to the whole body—and special senses associated with particular organs such as the eyes and ears. Somesthetic senses are broken down into exteroceptive (initiated by stimuli outside the body), proprioceptive (initiated within the body, determining the orientation of body parts relative to one another and the orientation of the body in space), and visceral (usually from internal organs and usually painful). Cetaceans, as far as is known, are subject to the familiar exteroceptive sensations. For example, captive and stranded animals respond to stimuli of touch, pain, and heat. Because precise assessment of the other somesthetic modalities (proprioceptive and visceral) is difficult, scientists have simply assumed their presence.

The special senses respond to stimuli registered by specialized organs or tissues. One way to quantify the presence of a special sense in an animal is to consider the organs involved.

Smell

The sense of smell can be defined as those sensations carried from nose to brain by the olfactory nerve. Toothed whales have lost the olfactory nerve, so by definition they are incapable of smelling. On the other hand, they do use "quasi-olfaction" (see below). Baleen whales have retained this nerve and have a reduced area for olfaction in the nasal passage, but this sense is active only while the animal is breathing at the surface.

Taste

Captive dolphins (family Delphinidae) commonly exercise food taste discrimination that is comparable to the human ability, in spite of the fact that the presence of taste buds in cetaceans has not been demonstrated. Regardless, dolphins have been shown to be sensitive to the standard four qualities of taste: sweet, salty, sour, and bitter. It has been established that the bottlenose dolphin (Tursiops truncatus) has a highly effective sense, called quasi-olfaction, operating through pits in the back of the tongue. This sense permits dolphins to experience what would be classified as smell, but quasi-olfaction does not involve the nasal passages.

Sight

Cetaceans have well-developed eyes and good vision. The popular notion that whales have reduced vision is probably based on the relative size of their eyes, but this assumption is functionally incorrect. Vision in both the water and the air has been experimentally evaluated in captive dolphins and found to be excellent. They have binocular vision over at least part of the visual field but are largely insensitive to colour. In one genus of river dolphin (Platanista of the muddy Ganges and Indus rivers), the eyes are reduced to organs that can detect only the difference between light and dark. The external opening for the eye is a slit only 2–3 cm (about an inch) long.

Hearing

Whales and dolphins have long been known to possess an acute sense of hearing. When approaching whales, whalers muffled their oars to prevent the animals from hearing them. Research done with captive animals in the 1950s quantitatively demonstrated that dolphins both produce and are sensitive to sounds into the ultrasonic range. Dolphins and porpoises were found to have the ability to derive information about their environment by listening to echoes of sounds that they have produced (echolocation). The amount of information obtained by an echolocating dolphin is similar to that obtained with the eyes of a sighted human.

The sound sensitivity of dolphins falls off near the bottom of the human acoustic spectrum (40–50 hertz), but this is the beginning of the range used by the large baleen whales. Fin whales (Balaenoptera physalus) and blue whales have been recorded producing subsonic sounds around 10 hertz and are capable of producing extremely loud noises at those frequencies. The strength of these vocalizations enabled one blue whale to be followed by fixed hydrophone arrays on the ocean bottom for 43 days over a course of 2,700 km (1,700 miles).

Magnetic sensitivity

Much interest has been shown in various animals’ ability to sense the Earth’s magnetic field. It has been demonstrated that birds and fish use magnetoreception in migration, and theories to explain why cetaceans beach themselves in mass strandings (see below) have included magnetic detection. Although magnetite has been found in some skulls of the common dolphin (Delphinus delphis), it has not been found in other specimens of the same species, and no conclusive data indicate its biological use.

Natural history

Life history

Distribution and migration

Cetaceans are distributed in all the world’s oceans from the far polar reaches to the Equator. They concentrate in areas of increased biological productivity, such as upwellings where there is an abundant supply of food. Some species are coastal, and some are pelagic, dwelling farther offshore. The centres of the ocean basins appear not to have any concentrations of whales or dolphins. Some small cetaceans are distributed in major river systems, particularly river dolphins of the family Platanistidae. Members of this family are found in the Amazon, Orinoco, La Plata, Yangtze, Ganges, and Indus rivers and surrounding drainage waters. Members of other families, particularly the Delphinidae and Phocoenidae, spend part of their time in fresh water.

As a rule, large whales have north and south seasonal migrations, spending summers in high latitudes near the poles, where there is an abundant food supply, and moving toward the Equator in the fall to breed. Some populations of these species, however, reside in one locality all year. One of the greatest migrations is undertaken by the California population of the gray whale, which summers in the Bering and Chukchi seas of the Arctic and winters in lagoons off the coast of Baja California—a journey of 5,000 km (about 3,000 miles) each way.

Migratory whales usually do not cross the Equator, and that has led to the development of genetically separate populations in the north and south ocean basins. However, one humpback whale (Megaptera novaeangliae) was photographically identified near the Antarctic Peninsula and was later sighted on the coast of Colombia, having covered at least 8,334 km in both the South Atlantic and North Atlantic ocean basins. Some sperm whales (Physeter catodon) sexually segregate on their migrations, with larger and older males going much farther in a polar direction during the summer.

Diet

All cetaceans are carnivores and do not consume plants or algae as food. The large baleen whales eat schooling organisms that range in length from minute drifting mollusks, copepods (1 cm or less), krill (1–5 cm), and small fish and squid up to about 40 cm. All these are consumed by whales in vast quantities with each concentrated mouthful. Certain smaller baleen whales, such as the minke whale (Balaenoptera acutorostrata), also pursue individual fish up to 1 metre long. Toothed whales, which range in length from about 1 metre for the finless porpoise porpoises (Neophocoena phocaenoidesand N. asiaeorientalis) to 20 metres for the sperm whale, eat an enormous variety of prey ranging from small shrimp, fish, and squid to bluefin tuna (3 metres long) and giant squid.

Cetaceans void their solid digestive waste products as pastelike feces, enabling the retention of intestinal water. Liquids are excreted in the urine. Biologists do not entirely understand the function of ambergris, which seems to be a normal digestive secretion of the sperm whale.

Reproduction

Cetacean breeding is seasonal, usually in the winter, and females normally calve once every two years. As mammals, they reproduce by internal fertilization. The testes and penis of the male are internal, but the penis is capable of being extended and introduced to the female during mating. After the female’s egg has been fertilized, she carries the fetus for about a year, although some toothed whales have gestations of up to 18 months. Cetaceans give birth tail first, opposite of most terrestrial mammals. The mothers produce extremely rich milk for their young—50 percent fat is common. The mammary glands are paired and located at the lower abdomen, just forward of the anal-genital slits. One calf is born (multiple fetuses have been found, but never live twins), which is weaned at six months to a year but continues to grow rapidly until 5 to 10 years of age. The largest whale, the blue whale, is born with a length of about 7.3 metres and a weight of about 3 tons; it grows an average of 0.3 metre per week and gains weight at a rate of 90 kg (nearly 200 pounds) per day.

Age and growth

Sexual maturity occurs at an age of about 6–10 years and is defined in cetaceans as the age at which the females start ovulating and are capable of becoming pregnant. Generally, male cetaceans reach a slightly greater size than females, but there are many exceptions; female baleen whales and some of the beaked whales, river dolphins, and porpoises tend to be slightly larger than males of their species. Male sperm whales, on the other hand, are on average 50 percent larger than females. Physical maturity is defined specifically in cetaceans as the point when all of the vertebrae stop growing, which occurs at an age of about 8–25 years. Historical estimates of whale longevity are limited because of difficulty measuring age in older whales. Modern techniques have determined some fin whales to be 100 years old, some humpbacks 96, and some blue whales 90. But the longest-lived cetacean by far is the bowhead, a right whale that can survive for more than 200 years.

Abundance

Counting animals that can be spread over a wide areas of the world’s oceans and are visible for only a few seconds while they breathe is extremely difficult and expensive. The largest populations existed in the oceans around Antarctica, where harsh and remote conditions make biological research difficult and infrequent. The abundance of cetaceans is thus hard to estimate accurately, but whale populations have varied over the years, depending largely upon human activities.

Biologists estimate that there were 228,000 blue whales and 548,000 fin whales in the world’s oceans when modern whaling began in the early 20th century. At the beginning of the 21st century, there were an estimated 14,000 blue whales and 120,000 fin whales left. California gray whales were thought to number 20,000 in 1847, then were hunted until they were thought to be extinct in the 1920s. Since then the species has recovered under protective legislation, and its population has been estimated to be more than 26,000.

Until the early 21st century, the only cetacean population to be completely exterminated was the Atlantic gray whale, which was gone in the early 1700s; however, the baiji, or Chinese river dolphin (Lipotes vexillifer), a species restricted to the Yangtze River (Chang Jiang), was widely believed to be extinct. In addition, some smaller cetacean species with limited distributions—such as the Gulf of California porpoise (Phocoena sinus), Indus susu (Platanista gangetica minor), and Ganges susu (P. g. Platanista gangetica)—could be in immediate danger of becoming extinct. (Some sources consider P. minor and P. gangetica as subspecies rather than separate species.) Furthermore, many other cetacean species, such as the gray whale and the northern right whale (Eubalaena glacialis), continue to be menaced by ship collisions, pollution, entanglement in commercial fishing equipment, and illegal hunting.

Diseases and parasitism

Cetaceans can suffer from many of the same diseases and parasites that afflict humans and other mammals: cancer, arthritis, pneumonia, lungworms, tapeworms, and roundworms, to name just a few. In the 1980s various dolphin species experienced epidemics of a morbillivirus, a disease similar to distemper and measles.

Of the parasites and commensal organisms, some are also found on fish and marine turtles, and others are specific to cetaceans. Commensal barnacles are most visible on humpbacks and gray whales, although they occur to a lesser extent on many other baleen and toothed whales. Xenobalanus globicipitis, a unique type of small pseudo-stalked barnacle, occurs on the appendages of cetaceans, including the common bottlenose dolphin. Stalked barnacles can also occur on exposed teeth and can be particularly striking on the tusks of beaked whales.

Different types of commensal or parasitic crustaceans inhabit whales. There is a small commensal copepod, Balaenophilus, that eats the algae on the baleen of some rorqual species. A specific family of amphipods (Cyamidae) called whale lice routinely infest right, humpback, and gray whales and also occur opportunistically on most species of baleen and toothed whales, particularly around wounds. They appear to eat sloughed skin. Another crustacean is clearly parasitic and is a member of the caligoid copepod genus Pennella. It is commonly about 2–10 cm long and lives with its body buried in the blubber or skin of cetaceans and fishes.

Small (4–7 cm) circular scars left in the skin of whales, dolphins, and fish were a mystery during the early 20th century. Explorers reported that the scars were created by an unknown organism that they called the “DWB” or “demon whale biter,” which cleanly removed hemispheric chunks of blubber as though extracting them with a razor-sharp scoop. The creature responsible was finally identified in the 1950s as a grazing predator, the cookie-cutter, or cigar, shark (genus Isistius).

Locomotion

Swimming

As described in the section General features, cetaceans Cetaceans swim by using vertical tail movements that drive the horizontal flukes up and down, powered by the long epaxial and hypaxial muscles that lie along the spine. The tail flexes through a point between the dorsal fin and the anus, while the thorax and abdomen are relatively inflexible. The body itself acts like a spring to propel the animal through the water with minimal energy.

Much was written about the speeds of cetaceans in the mid-20th century. It seemed that cetaceans could exceed the speed at which turbulence would make locomotion energetically very expensive. However, the swimming-speed figures were estimates that turned out to be very high. Further investigation found that, regardless of size, the cruising speed of most cetaceans is about 2 metres per second (about 7 km, or 4 miles, per hour). A combination of biomechanical and hydrodynamic factors make this an efficient speed at which to travel. Maximum speeds, however, vary greatly between species.

Common dolphins (genus Delphinus) have been observed keeping pace with boats for a considerable period of time at 36 km/hr (kilometres per hour). Researchers trained Pacific bottlenose dolphins (genus Tursiops) to swim in an open-water environment, thus removing the spatial limitations of a pool while conserving experimental controls. They found that the dolphins could sprint at 29.9 km/hr for 7.5 seconds and could maintain speed at 21.9 km/hr for 50 seconds. When dolphins ride a bow wave, they coast at the speed of the ship while expending very little energy (see below). Fin and blue whales can swim fast enough that a boat must travel in excess of 30 km/hr to catch up to them, and they can maintain speeds of 33–37 km/hr for periods of up to 10–15 minutes. Sonar records indicate that fin whales can sprint at 48 km/hr. Right, humpback, and gray whales, however, can seldom swim faster than 9 km/hr. Sperm whales can cruise at 7.5 km/hr and swim up to 36 km/hr in spurts. The fastest cetacean appears to be the sei whale (Balaenoptera borealis), recorded moving at speeds up to 65 km/hr along the ocean surface.

Breathing and diving

Cetaceans surface periodically to breathe, and the intervals between breaths vary depending on what the animal is doing. Intervals may range from about 20 seconds for dolphins that are actively swimming to 5–10 minutes for a resting blue whale. A common breathing pattern in large whales is to breathe every 20 seconds for 8–10 breaths and then dive for about 10–15 minutes. Most whales stay in the upper 100 metres of water. Deep-diving whales—such as the sperm whale, which has been recorded diving to depths of 1 km—may stay down for an hour. The longest recorded dive is that of a harpooned bottlenose whale (Hyperoodon ampullatus) that dived for two hours, surfaced, and then dived again. Patterns of locomotion and breathing are very important to whale watchers identifying whales at a distance, as different species show different blow heights and shapes. Right whales, for instance, have an unequal inclination to their two nasal passages, so their blows appear in pairs. Humpbacks and gray whales have blows that appear low and wide (bushy), and sperm whales have a bushy blow that is angled to the left and forward.

Leaping and wave riding

Small cetaceans “porpoise” when they are swimming rapidly; that is, they rise out of the water in a low leap that keeps the head clear of the water for breathing. Spinner dolphins (Stenella longirostris) frequently leap out of the water while spinning on their long axis, hence their common name. Trained porpoises and dolphins can leap straight up as high as six metres. Leaping is very rare in large whales, but some rorquals (genus Balaenoptera) have been photographed jumping clear of the water.

Many small cetaceans play around moving boats, where they bow-ride, taking advantage of their ability to bodysurf and essentially enjoying the free ride in the bow wave created by the vessel. They also practice this behaviour around large whales that are swimming fast enough to produce a bow wave.

Behaviour

Social behaviour

All cetaceans are social to some extent. The minimum group of mother and calf is commonly expanded to a nuclear family or a group of closely related individuals. A group of cetaceans that normally feed and travel together is called by various names: school, herd, pod, or gam. It is often difficult to define or measure, as its members can be spread over kilometres of ocean but still be in contact with one another. Sometimes these schools coalesce into even bigger groups of more than 1,000. Groups of whales can persist for many years, and studies of coastal dolphins have shown long-term association of dolphins with their mothers. Groups (particularly of small toothed whales) frequently associate with other cetacean species. For example, associations between pilot whales and bottlenose dolphins have been observed, as have associations between common dolphins and fin whales.

Play is a common behaviour, especially among young animals. Play allows individuals to practice and perfect behaviour patterns, such as aggression, that will be socially useful later in life; a significant portion of play is sexually oriented. Captive dolphins have also been observed playing with fish, birds, and turtles.

Many cetaceans exhibit epimeletic behaviour, in which healthy animals take care of another animal that has become temporarily incapacitated. This is evident when a wounded or sick whale is supported by others or in cases when a dolphin (usually the mother) pushes a dead calf around.

Cetaceans show fright by fleeing from a situation or by bunching up and "milling." The former response has been utilized by fishermen, who drive a whale or school of dolphins into a situation where they can kill it. Milling has been seen in dolphin schools driven into an enclosure or caught in a net; the animals move in a circle or eddying mass, and at the height of this reaction they stop swimming, sink, and die.

Aggression and defense

Aggression is common among cetaceans and is seen in normal herd behaviour and feeding. One form of aggression helps to establish social hierarchy: the dominant animal nips the less-dominant animal, which produces the tooth scars seen on every adult in the dolphin family (Delphinidae). Mating behaviour also involves biting, as one of the ways males compete for females is by biting and raking the teeth over another male. Adult male beaked whales (family Ziphiidae) have very densely ossified rostra (beaks) used as weapons in combat for females. Another more dangerous means of aggression is head butting. Cetaceans can ram their heads into other individuals and kill them. This has been seen in captivity and in aggressive behaviours toward other species such as sharks and accounts for many of the broken ribs and vertebrae seen in stranded animals.

Normally, aggression is associated with members of the same species or as a defense response to predation from other species. Although cetaceans can defend themselves by utilizing the behaviours of intraspecific aggression (biting, ramming, and butting), the primary weapon that cetaceans have for self-defense is the tail. Cornered whales slash sideways with their flukes and can incapacitate a bigger whale or a boat. Head butting as a form of defense was immortalized in the 1820 sinking of the whaling ship Essex by a sperm whale.

Courtship and mating

Sexual behaviour starts early in cetaceans. Young dolphins engage in exploratory sexual behaviour involving their mothers and other members of the school. Self-stimulation is common in both sexes. Male cetaceans perhaps use their penises as a manipulation organ in much the same way that people use their hands. This exploratory behaviour gradually becomes courtship and mating behaviour.

Courtship involves physical and acoustic displays, such as the elaborate songs of male humpbacks, and leads to contact with the flippers and other parts of the body. Successful courtship culminates in mating. Copulation is relatively brief in cetaceans. It can be secretive, or it can be boisterous as in the mating displays of right and gray whales, when a number of males attempt to mate with a single female.

Feeding

Cetaceans hunt as individuals or in schools. When hunting in schools, dolphins or whales herd their prey in order to concentrate a large volume before eating. Hunting alone is preferred where prey is more scattered.

Before they swallow their food, toothed whales disable it; biologists think that some can stun their prey by emitting a high-energy burst of sound. Normally, cetaceans eat animals that can be swallowed intact, as their teeth are shaped for holding, not chewing. If, however, the prey is too large to swallow in one bite, it is ripped into chunks. Killer whales (Orca orcinus) have been seen to grab seals and shake them in the air so hard that the bodies come apart.

Baleen whales also herd their prey like toothed whales, but they engulf it in either of two feeding methods: gulp or skim. In gulp feeding, the whale opens its mouth to take in a huge mouthful of water, closes its mouth, strains the water out through the baleen apparatus along the sides of the mouth, and swallows its prey. Gulp feeding is common in rorquals, which have ventral grooves that stretch to enlarge the oral cavity. One of the rorquals, the sei whale, as well as the nonrorqual baleen whales (right, bowhead, pygmy right, and gray), skim-feed by locating a concentration of zooplankton prey and swimming through it with the mouth open. Skimming may last up to several minutes until the whales close their mouths to swallow what they have filtered from the water.

Sleep

Breathing is a conscious activity in cetaceans; they must consciously breathe, or they will drown. Therefore, they cannot enter into what humans understand as unconscious sleep; instead, they have periods of little activity but not total inactivity. Studies of dolphins have revealed that they shut down half of their brain during sleep. The other half of the brain stays awake to signal when to rise to the surface to breathe and to watch for predators and obstacles. Large whales appear to surface-sleep. Floating horizontally just below the water’s surface, they move their flukes periodically to rise above the water for a breath.

Intelligence

Although several cetaceans are easily trained and much has been theorized about the possible intelligence of whales and dolphins, little is known for certain. Some researchers equate brain size with intelligence, reasoning that cetaceans should have the capacity for intelligence because they have relatively large brains. The human brain averages about 1.2 kg, the bottlenose dolphin brain about 1.8 kg. The largest cetacean brain recorded was a sperm whale’s, weighing 9.2 kg. However, cetaceans may use their increased brain weight for processing acoustic information. In any event, it seems unproductive to compare species with which it is difficult even to communicate until a definition of nonhuman intelligence has been refined.

Stranding

Stranding is a phenomenon that has long fascinated people, and there is fossil evidence of mass strandings from before humans evolved. Many stranded cetaceans are found already dead, and it is not known if they were alive and conscious when they stranded themselves. When a whale or dolphin dies offshore, it usually sinks; if the water is shallow enough to permit decomposition gases to form, it will float ashore, so some stranding represents normal mortality. If infection or some other factor interferes with a cetacean’s ability to navigate, it could come ashore while still alive—though most cetaceans have difficulty out of water and usually die. These cases are known (alive or dead) as single strandings. Sometimes up to several hundred toothed whales swim ashore, and this phenomenon is known as a mass stranding.

There are no records of the mass stranding of baleen whales; all such events have involved only toothed whales that normally live offshore and may not be familiar with physical borders. Perhaps not realizing that the ocean has a bottom and sides, they may somehow enter shallow water and find themselves unable to deal with the strange environment. Because they are also members of extremely social species and may be kept together by group ties, they may have even greater difficulty extricating themselves. In any case, biologists are beginning to realize that cetaceans are behaviorally complex enough that a simple blanket explanation of mass stranding is not likely to be valid. Biologists have tried to attribute mass stranding to a number of causes:

1. Something wrong with the leader of a group2. Epidemic disease 3. Getting lost in pursuit of prey4. Parasitic infestation that affects the hearing5. Following migratory routes laid down by remote ancestors6. Magnetic anomalies that lead the school astray7. Behavioral reversion to a period when cetacean ancestors were terrestrial and land was a haven8. Fright reaction to predators9. Failure of echolocation signals to work properly in shallow water10. Overpopulation11. Suicide

Sound production and communication

All cetaceans produce sound, some more extensively than others, and they primarily use the larynx for this purpose. At one time it was argued that the cetacean larynx was incapable of generating sound because it does not have vocal cords. However, vocal cords are restricted mainly to primates; dogs and cats, for example, have vocal folds, and both baleen and toothed whales possess structures that modify sound. Baleen whales have laryngeal pouches, and toothed whales have accessory air sacs and fat bodies in their noses. In addition, toothed whales can generate high-frequency sounds in their nasal passages.

Cetacean sounds can be roughly divided into communication signals and echolocation signals. Communication does not necessarily imply language, and it can simply be one-way, as when one dolphin knows another is present because the second dolphin is vocalizing. Echolocation, which involves generating certain sounds and listening to the echoes of those sounds, has been recognized in toothed whales but not baleen whales. Toothed whales use extremely high frequencies, on the order of 150 kilohertz, for refining spatial resolution from their echoes. They are capable of “seeing” into and through most soft objects such as other dolphins, though the effectiveness of toothed whale echolocation drops off at distances greater than about 100 metres. To produce such high frequencies, toothed whales possess modified tissues associated with the blowhole on the right side of the head; the left side is not modified, and the result is skull asymmetry. This condition is extreme in sperm whales, which is not surprising, as most of the head is involved with sound production. The head of a 16-metre adult male sperm whale is about 6 metres long, 3 metres high, and 3 metres wide, a mass of tissue that can weigh about 20 tons. The bulk of it is occupied by the spermaceti organ and a fatty (adipose) cushion, both of which somehow function in the emission of sound for echolocation and were known by whalers as the “case” and the “junk,” respectively. The junk of the sperm whale is the fatty structure found in the forehead of other toothed whales and known by whalers as the “melon” because of its pale yellow colour and uniform consistency. Baleen whales generate sounds at frequencies that are audible to humans (sonic) or below that range (subsonic). Some of their vocalizations are very loud; biologists have recorded extremely low sounds (12.5–200 hertz) from a blue whale and have claimed they are the loudest sounds known from any animal. The songs that humpbacks use for courtship were brought to public awareness in 1971. Baleen whales (mysticetes) use calls like these for communication and possibly for low-frequency long-range echolocation in orientation and navigation. Their low-frequency sounds are powerful enough that mysticetes might be able to communicate across entire ocean basins.

Paleontology and classification

Paleontology

Cetaceans are distant descendants of a group of poorly defined mammals known as condylarths. There is debate as to whether the first cetaceans (archaeocetes) descended from an extinct group of large carnivores called mesonychids or from a group of hoofed herbivores (artiodactyls). The earliest archaeocetes were huge dolphinlike creatures 6 to 10 metres long. Basilosaurus (Zeuglodon) was an unusual genus that was up to 34 metres long, but it apparently gave rise to no descendant groups.

As the fossil record becomes more complete, the pattern will emerge as to which condylarth is ancestral to archaeocetes and which archaeocete is ancestral to living cetaceans. The first fossil cetacean, Pakicetus, is known from the Early Eocene Epoch (55.8 million to 49 million years ago) in Pakistan. It has recently become clear that archaeocetes rapidly diversified during the Eocene, and at least five now-extinct families are recognized. One subfamily of the Basilosauridae, the Dorudontinae, is thought to have given rise to both living suborders of cetaceans (baleen whales and toothed whales) sometime during the Late Oligocene Epoch, about 25 million years ago.

The first baleen whales had wide, flat skulls bordered by a reduced number of teeth in the archaeocete pattern. The roof of the mouth widened between these borders, and grooves for blood vessels supplying the emerging baleen are seen inside the tooth rows. By the middle of the Miocene Epoch (some 16 million to 11 million years ago), there were several families of baleen whales, including the right whales and rorquals. The Miocene was the epoch during which modern ocean circulation began; regional areas of upwelling and increased productivity developed, setting the stage for the evolution of large whales with seasonal migratory distribution. At the same time, the modern toothed whales began to emerge, developing into nine families during this period; four of these have since become extinct. Sperm whales were among the first toothed whales and were present during the Middle Miocene as large and well-defined as they are now. From the fossil record it is evident that today’s cetacean biodiversity has decreased markedly since the Miocene.

Classification

Though there is some consensus among taxonomists that Cetacea should be treated as one order (as they are in this article), others believe they are actually two or three. This depends on the evaluation of the degree of shared ancestry, which remains controversial. It is possible that certain protocetids, or “pre-whales,” could have given rise to both modern groups of whales, which is why some authorities prefer a single order. However, the absence of intermediate fossils linking baleen whales with toothed forms supports the use of separate orders. Resolution of this problem awaits the discovery of relevant fossils.

Further disagreement occurs at many points below the ordinal level. Although there is no doubt that any recent cetacean is either toothed (suborder Odontoceti) or baleen (suborder Mysticeti), the relationships of many genera are in doubt. For example, the long-snouted dolphins are classified by some authorities as a separate family, Stenidae, rather than family Delphinidae. A similar situation exists for the porpoises (family Phocoenidae). At the species level there is uncertainty about the specific or subspecific status of many populations.

It must be borne in mind that all classifications are, to an extent, artificial. Over time one species merges with another, and some classification issues actually get more complex as the fossil record improves. Current cetacean classification is the result of an improving fossil record that reveals more taxa near the origins of the three established suborders (the archaeocetes, mysticetes, and odontocetes). It has become apparent that a major diversification was associated with each major adaptive branch, and species are currently being shuttled back and forth between the latest representatives of one family and the earliest representatives of its descendant family.

Although species relationships have historically been based on morphological (anatomic) characteristics, biologists have begun comparing DNA sequences of different cetaceans and thereby causing morphologists to reexamine their taxonomic data. Chromosome count (karyotype) can be quite variable between related mammals, but it is remarkably stable among cetaceans. All baleen whales and most toothed whales have 44 chromosomes, and sperm whales and beaked whales have 42. Techniques such as DNA sequencing have provided different ways of evaluating the relationships of species. Applications of these and older techniques should provide a clearer understanding of the evolutionary history of Cetacea.

Annotated taxonomy

The three cetacean suborders (two living, one fossil) recognized below share the same basic body plan but differ in their degree of specialization. Suborders and families are separated primarily on the basis of the following characteristics: tooth structure, number, and degree of differentiation; skull modifications, especially the position of the nostrils, degree of telescoping of the whole skull, modifications to the inner and middle ear, and extent of joining of the two halves of the jaw; and degree of modification of the pelvic girdle (archaeocetes only). The classification presented here is based on research by cetologists F.C. Fraser, R. Kellogg, and a number of other modern authorities. Groups marked with a dagger (†) are extinct and known only from fossils.

Order Cetacea (whales, dolphins, and porpoises)81 species in 2 suborders. Aquatic mammals with forelimbs modified into flippers, hind limbs lacking; pelvic girdle vestigial and not attached to vertebral column; tail laterally flattened and extended into horizontal flukes, supported by fibrous connective tissue. External nares present as blowholes at top of head (except in sperm whales and archaeocetes); cranium telescoped variably, with elongated rostrum. 1 fossil suborder. Suborder Odontoceti (toothed whales)69 species in 6 families. Worldwide, marine, a few in fresh water. Teeth relatively uniform (homodont) and numerous (up to 300), occasionally reduced. Skull asymmetrical; nostrils on top of head (except in Physeteridae) with single external opening (blowhole). Mandibles fused anteriorly. Several pairs of ribs joining sternum, which is fused into a single bone only in the adult. 9 extinct families. Upper Eocene to Holocene.Family Delphinidae (dolphins and killer whales)32 species in 12 genera. Characteristics variable. Some neck vertebrae fused; dorsal fin and well-defined beak present or absent. None with expanded tooth crowns. Length 1.7 to about 9 metres. Worldwide, marine and fresh water. About 35 fossil genera, Lower Miocene to Holocene. Family Ziphiidae (beaked whales)21 species in 6 genera. Previously called Hyperoodontidae. Some neck vertebrae fused; functional teeth greatly reduced in number to 1 or 2 pairs in lower jaw (except for total 100 teeth in Tasmacetus); vestigial (nonfunctional) teeth not uncommon. Pair of longitudinal grooves in throat region. Dorsal fin slightly more than one-third of way back along body. Tail usually without middle notch. Flippers small. Worldwide, marine. Length to about 12 metres. About 14 fossil genera. Lower Miocene to Holocene. Family Phocoenidae (porpoises)6 7 species in 3 genera. Some neck vertebrae fused; well-defined beak absent; teeth with expanded spade-shaped crowns. Dorsal fin triangular, when present. Adult length in living species to about 2 metres. Virtually worldwide, marine, 1 2 species living in some rivers in China and Japanthe coastal waters and rivers along the Indian Ocean and western Pacific Ocean. 3 fossil genera. Upper Miocene to Holocene. Family Platanistidae (Indian river dolphins)5 2 species in 4 genera1 genus. Snout long and slender; neck vertebrae all free. Flippers short and broad, sternum well developed. Adult length 1.5–2.5 metres. Found in the Ganges, Indus, and Brahmaputra rivers of India; the Indian subcontinent. About 10 fossil genera. Lower Miocene to Holocene. Family Iniidae (river dolphins)3 species in 2 genera. Snout long and slender; neck vertebrae all free. Flippers short and broad, sternum well developed. Adult length 1.5–2.5 metres. Found in the La Plata, Amazon, and Orinoco rivers of South America; and . A third genus, Lipotes, from Dongting Lake and the adjacent Yangtze River of China. About 10 fossil genera. , is thought to have become extinct in the early 21st century. Lower Miocene to Holocene.Family Physeteridae (sperm whales)3 species in 2 genera. Some neck vertebrae fused; head disproportionately large, with bulbous, squared snout; mouth narrow and ventral; lower teeth total 40–52 in Physeter, 16–32 in Kogia; upper teeth vestigial, smaller, varying in number. Blowhole a single asymmetrical opening on the front left tip of the snout. Length to 19 metres (Physeter) and to 3.4 metres (Kogia) in Holocene genera. Tropical and temperate oceans, adult males to polar waters. 20 fossil genera. Lower Miocene to Holocene.Family Monodontidae (narwhal and beluga)2 species in 2 genera. Dorsal fin lacking. Neck vertebrae free. Flippers broad, rounded at tips. Teeth reduced to 8 or 10 in Delphinapterus (beluga); all teeth vestigial in Monodon (narwhal) except for 1 left tooth of male, which grows into a long, straight tusk extending in front of the animal. Length to about 5 metres. Arctic Ocean. Pleistocene to Holocene. †Family Agorophiidae1 genus. Lower Oligocene of North America. †Family SqualodontidaeAt least 12 genera. Upper Oligocene to Late Miocene. Europe, North and South America, Australia, and New Zealand. †Family Squalodelphidae3 genera. Lower Miocene. Europe and South America. †Family Dalpiazinidae1 genus. Early Miocene. Europe. †Family Kentriodontidae11 genera. Upper Oligocene to Late Miocene. South America, Europe, Asia, and North America. †Family AlbireodontidaeUpper Miocene to Pliocene. North America. †Families Eurhinodelphidae, Hemisyntrachelidae, and AcrodelphidaeAbout 12 genera of Miocene toothed whales similar to today’s dolphins. All continents. Suborder Mysticeti (baleen whales)12 species in 4 families. Teeth lacking in adult, numerous but vestigial in embryo; baleen present. Lower jaw large, bowed outward, loosely united in front. Skull symmetrical; relatively well-developed nasal bones; blowholes are paired slits. Family Balaenopteridae (rorquals and humpback whale)8 species in 2 genera. Skull broader and less arched than in Balaenidae; baleen plates shorter, broader, less flexible; neck vertebrae not fused. Dorsal fin present; flippers narrow. Conspicuous longitudinal grooves on throat. Length 10 to perhaps 33.6 metres; blue whale (Balaenoptera musculus) is the heaviest animal that has ever lived. Family occurs in all oceans and, at various times, from Equator to poles. 8 fossil genera. Upper Miocene to present. Family Eschrichtiidae (gray whale)1 species. Formerly Rhachianectidae. Head less than one-quarter of total length; neck vertebrae not fused. Dorsal fin lacking. Flippers with 4 internal digits. Length to about 15 metres. Found along both coasts of the North Pacific Ocean. Exterminated in the Atlantic Ocean in the 18th century. Pleistocene to Holocene.Family Balaenidae (right whales)4 species in 2 genera. Skull significantly arched; neck vertebrae fused; baleen long, narrow, flexible strips. Ventral grooves on throat lacking. Length 6–20 metres. Polar and subpolar oceans. 4 fossil genera. Lower Pliocene to present. Family Neobalaenidae (pygmy right whale)1 species. Skull slightly arched; neck vertebrae fused; lower jaw broad; baleen long and narrow; ventral grooves lacking; dorsal fin present; proximal portion of ribs extremely widened; lumbar vertebrae reduced to 2. Length 6 metres. Southern Hemisphere in temperate waters. †Family Aetiocetidae1 genus, possibly 2. Upper Oligocene. Toothed but with symmetrical skull and other typical mysticete features. North America. †Family Mammolodontidae1 genus. Upper Oligocene? Lower Miocene? Australia. †Family Kekenodontidae 2 genera. Upper Oligocene. Europe and New Zealand.†Family Patriocetidae4 genera. Upper Eocene to Upper Oligocene. Europe and North America. †Family Llanocetidae1 genus. Lower Oligocene or Upper Eocene. Antarctica. †Family Cetotheriidae (cetotheres)About 30 genera. Middle Oligocene to Lower Pliocene. North and South America and Europe. †Suborder Archaeoceti (archaeocetes, or zeuglodonts)At least 24 genera. Anterior and posterior teeth differentiated; total teeth not exceeding 44, the basic number in terrestrial placental mammals. Nostrils positioned toward the top of the head to varying degrees. Lower Eocene to Middle Miocene. †Family Pakicetidae 3 genera. Early Eocene. India and Pakistan. †Family Ambulocetidae2 genera. Early to Middle Eocene. Pakistan. †Family Remingtonicetidae4 genera. Middle Eocene. Pakistan. †Family Protocetidae9 genera. Early to Middle Eocene. Europe, Africa, and possibly North America. †Family Dorudontidae5 genera. Middle to Upper Eocene. Europe, Africa, and North America. †Family Basilosauridae1 genus. Middle to Upper Eocene. Europe, Africa, and North America.